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Journal of Bacteriology, July 2004, p. 4262-4275, Vol. 186, No. 13
0021-9193/04/$08.00+0     DOI: 10.1128/JB.186.13.4262-4275.2004
Copyright © 2004, American Society for Microbiology. All Rights Reserved.

Autoinduction of Bacillus subtilis phoPR Operon Transcription Results from Enhanced Transcription from E{sigma}A- and E{sigma}E-Responsive Promoters by Phosphorylated PhoP

Salbi Paul, Stephanie Birkey,{dagger}, Wei Liu,{ddagger}, and F. Marion Hulett*

Laboratory for Molecular Biology, Department of Biological Sciences, University of Illinois at Chicago, Chicago, Illinois 60607

Received 3 February 2004/ Accepted 1 April 2004

The phoPR operon encodes a response regulator, PhoP, and a histidine kinase, PhoR, which activate or repress genes of the Bacillus subtilis Pho regulon in response to an extracellular phosphate deficiency. Induction of phoPR upon phosphate starvation required activity of both PhoP and PhoR, suggesting autoregulation of the operon, a suggestion that is supported here by PhoP footprinting on the phoPR promoter. Primer extension analyses, using RNA from JH642 or isogenic sigE or sigB mutants isolated at different stages of growth and/or under different growth conditions, suggested that expression of the phoPR operon represents the sum of five promoters, each responding to a specific growth phase and environmental controls. The temporal expression of the phoPR promoters was investigated using in vitro transcription assays with RNA polymerase holoenzyme isolated at different stages of Pho induction, from JH642 or isogenic sigE or sigB mutants. In vitro transcription studies using reconstituted E{sigma}A, E{sigma}B, and E{sigma}E holoenzymes identified PA4 and PA3 as E{sigma}A promoters and PE2 as an E{sigma}E promoter. Phosphorylated PhoP (PhoP~P) enhanced transcription from each of these promoters. E{sigma}B was sufficient for in vitro transcription of the PB1 promoter. P5 was active only in a sigB mutant strain. These studies are the first to report a role for PhoP~P in activation of promoters that also have activity in the absence of Pho regulon induction and an activation role for PhoP~P at an E{sigma}E promoter. Information concerning PB1 and P5 creates a basis for further exploration of the regulatory coordination or overlap of the PhoPR and SigB regulons during phosphate starvation.

* Corresponding author. Mailing address: Laboratory for Molecular Biology, Department of Biological Sciences, University of Illinois at Chicago, 900 S. Ashland Ave. (M/C 567), Chicago, IL 60607. Phone: (312) 996-5460. Fax: (312) 413-2691. E-mail: Hulett{at}uic.edu.

{dagger} Present address: NFRP/TSCRP Program, Congressionally Directed Medical Research Programs, Science Applications International Corp., Ft. Detrick, MD 21702.

{ddagger} Present address: Genencor International, Inc., Palo Alto, CA 94304.

Journal of Bacteriology, July 2004, p. 4262-4275, Vol. 186, No. 13
0021-9193/04/$08.00+0     DOI: 10.1128/JB.186.13.4262-4275.2004
Copyright © 2004, American Society for Microbiology. All Rights Reserved.



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